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The Multinational Coordinated Arabidopsis thaliana

Genome Research Project

Progress Report: Year Two


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Any opinions, findings, conclusions, or recommendations expressed in this report are those of
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Cover (Flower of Arabidopsis thaliana) is courtesy of Dr. Keith Roberts, John Innes Institute.
The Multinational Coordinated Arabidopsis thaliana Genome
Research Project

Progress Report: Year Two

The Multinational Science Steering Committee

1992
PREFACE

In 1990, an ad hoc committee composed of nine scientists from the United States,
Europe, Japan and Australia prepared a report entitled "Long-range Plan for the Multinational
Coordinated Arabidopsis thaliana Genome Research Project". This publication (NSF 90-80)
outlined a plan for international cooperation in understanding the structure and function of the
genome of the higher plant Arabidopsis thaliana. The Committee assumed responsibility in six
areas: (1) coordinate programmatic aspects of the Arabidopsis genome project; (2) communicate
with the informatics and biological resource centers that were to be established; (3) monitor and
summarize progress of scientific activities of participating laboratories; (4) serve as a liaison to
the broader plant biology community; (5) identify needs and opportunities of the Arabidopsis
research community and communicate these needs to the funding agencies of participating
nations; and (6) periodically update the long-range plan.
This report of the Multinational Science Steering Committee is intended to summarize
progress made during the past year and to foster communication among those scientists with a
primary research interest in Arabidopsis as well as with other plant scientists and the general
scientific community. We also identify areas of present need for the international group of
funding agencies concerned with the project, and set forth a series of new goals for the coming
year. A previous report, describing progress made during the first year of the project is available
as publication NSF 91-60.
The information contained in this report was provided by members of the Multinational
Science Steering Committee and by others acknowledged below1. Sources include both the
published literature and less formal sources, including abstracts and presentations at symposia,
communications of the Arabidopsis electronic newsgroup, and personal communications. The
Committee outlined this report at Keystone, Colorado (USA) in April 1992. It is the nature of
any general progress report, representing the work of hundreds of scientists worldwide, that
some of the important experiments of the past year may have been overlooked or
misrepresented. Therefore, we ask not only that our colleagues overlook such shortcomings but
that they feel free to communicate their concerns and plans with their committee representatives,
so that future reports will be as accurate as possible.

The Multinational Science Steering Committee


Chairman: Marc van Montagu
Members: Caroline Dean, Richard Flavell, Howard Goodman, Maarten Koornneef,
Elliot Meyerowitz, Jim Peacock, Yoshiro Shimura, Chris Somerville

August 1992
I. Progress of the Previous Year

Because Arabidopsis is now widely used as an experimental organism, it has become

1 Among those to whom we are indebted for information used in the initial draft report, and for the subsequent
revisions are M. Anderson, F. Ausubel, A. Bent, M. Boguski, M. Caboche, M. Cherry, J. Chory, I. Crute, J. Dangl,
M. Delseny, D. Flanders, R. Gardner, R. Hangarter, E. Holub, D. Kristofferson, D. Meinke, K. Okada, S.
Pramanik, M. Schena, R. Scholl, P. Scolnick, P. Sijmons, S. Somerville, and R.F. Whittier.

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increasingly difficult to formulate a concise description of progress. The growth of information
about Arabidopsis, in particular, and plant biology, in general, is exploding. Much of the general
progress in this discipline is directly attributable to the fact that Arabidopsis has many attractive
features as an experimental organism. In addition, a substantial proportion of the visible
progress is attributable to the advantages of having a large number of scientists working on a
single organism where information, methods and materials can be freely exchanged. Several of
the initial objectives of the Arabidopsis genome project, which were designed to facilitate these
advantages, have been implemented during the past year and appear to have had the desired
effects. The results of these initiatives, and the projected effects of new initiatives are
summarized in the following sections under the major topics of genome analysis, technology
development, resource centers, informatics, human resource development and symposia. In
addition, various organized national research projects coordinated as the Multinational
Arabidopsis thaliana Genome Research project, are described.

1. Genome Analysis

In the original conception of the Arabidopsis Genome Research Project, a broad


approach to the dissection of the genome was envisioned. The rationale was that in order to
interpret the coding capacity, it was most desirable to characterize genes by all available criteria.
These included mutational analysis, the mapping of new mutations and genes, the mapping of
RFLP markers and the cloning and sequencing of genes on the basis of functional criteria, as
well as the eventual objective of obtaining the entire nucleotide sequence of the genome.
Progress on these fronts is summarized below under three headings: maps, mutations, and new
genes cloned and sequenced.

A. Maps

One of several promising genetic methods for the isolation of new genes is map based
cloning in which a gene is isolated solely on the basis of genetic map position. If a sufficiently
large number of RFLP markers were mapped, it would be possible to isolate virtually any gene
for which a mutation was known by probing a suitable genomic library with the RFLP markers
which closely flank the gene of interest, and isolating clones which carry both of the flanking
markers. It is, therefore, of substantial importance to have a high density, accurate genetic map
available for Arabidopsis. An international collaboration to integrate the two RFLP maps with a
total of approximately 300 markers, and the map of approximately 120 visible markers is
nearing completion. In addition, a new map based on randomly amplified polymorphic DNA
(RAPD) markers and recombinant inbred (RI) lines was published (Reiter et al., Proc. Natl.
Acad. Sci. USA 89,1477, 1992). This map contained 252 new markers which were mapped
relative to 60 previously mapped RFLP markers. The RI lines have been deposited with the
Nottingham and Ohio Resource Centers. Because they have been scored for a large number of
genetic markers, the availability of these lines will greatly facilitate the high resolution genetic
mapping of cloned genes.
Another project to add an additional 140 RFLP markers is nearing completion in Japan
(Mitsui Plant Biotechnology Research Institute, Tsukuba). These markers are being mapped
using another set of RI lines produced and characterized by C. Lister and C. Dean (Norwich,
U.K.) so there should be no impediment to the integration of these additional markers with

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existing maps. Hybridization of 30 markers from each of the two previously published maps
onto these RI lines has also improved the integration of the two maps.
Progress towards completion of a physical map continues on several fronts. A
collaborative effort by several U.S. laboratories and a British group has identified and mapped
yeast artificial chromosomes (YACs) covering approximately 33,000 kb or about one third of
the genome (Hwang et al., and Schmidt et al., Plant J. 1, 367 (1991)). The YAC libraries used
in these experiments are also being used to link up the approximately 750 cosmid contigs
representing the entire genome, which were produced by H. Goodman, B. Hauge and
collaborators (Boston, MA). A systematic effort on genome linking using YAC to contig
hybridization is underway in H. Goodman's laboratory (Boston, MA). To date, Goodman and
his colleagues have completed approximately 2,500 hybridizations (approximately four genome
equivalents of YAC clones) to the gridded cosmid filters containing the 1,920 cosmids and
approximately 500 hybridizations to the gridded cosmid filters with 480 cosmids. They have
written computer programs to transform the data into a form suitable for orienting with the
contig map, and are in the early stages of comparing the hybridization linkages with the
established contig map. The complete collection of 1,920 cosmids used on the filter grids has
been archived with the intention of making it available to the community.
As noted below, the widespread use of the YAC libraries for chromosome walking has
resulted in the construction of maps covering more than 1 million bp for at least four regions of
the genome. However, because of the presence of chimeric YACs and the frequent occurrence
of low copy number repetitive sequences in the Arabidopsis genome, constructing a physical
map by using ends of the YACs as hybridization probes has progressed slowly. Because of
increased interest in large-scale sequencing of Arabidopsis cDNAs (see below), the most
economical option may be to link up the YACs by hybridizing a large number of partially
sequenced cDNA clones to the YAC libraries. This would provide useful information about the
cDNA clone and would frequently provide information about overlap of YACs. Progress
toward a complete YAC map should also be greatly facilitated by the completion of a new YAC
library by J. Ecker and colleagues (Philadelphia, PA) which reportedly has significantly larger
inserts than previous libraries.
Several important initiatives, such as large-scale cDNA sequencing, and the development
of physical maps, are predicated on the concept that these methods will eventually intersect with
the results of research which is driven by specific biologically based questions. The probable
mechanism of integration is at the level of the genetic map. Thus, the utility of these initiatives
will ultimately depend upon the richness of the information anchored by mapped mutations. The
number of published newly mapped mutations increased by approximately 20% during the past
year. As an indication that significant expansion of the genetic map is pending, work is
currently underway in the Meinke laboratory (Stillwater, OK) to map 100 new embryo defective
mutations.
B. New Mutations Characterized

If the complete sequence of the Arabidopsis genome were available today, it would not
be possible to deduce the function of most of the genes because a relatively small proportion of
the genome has, as yet, been marked by informative mutations. Therefore, one of the highest
priorities for the genome project remains the development of an exhaustive collection of
characterized and mapped mutations. Although it has not been possible to catalog all of the
newly described mutations, progress on this front continues at an explosive pace. Indeed, one of

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the plant biology journals (Plant J.) has instituted a new section for short reports of new
mutations in a manner analogous to the short reports of new gene sequences which were used
before the advent of widespread and facile access to the nucleotide sequence databases.
Unfortunately, there appears to be a substantial lag between the isolation of a new mutation and
its appearance on the linkage map. It is hoped that the development of several new databases
(see below) will facilitate and accelerate the dissemination of this important information.
An example of recent progress in the identification of new mutations during the past year
has been the maturation of a broadly based thrust to develop well-defined host/pathogen systems
of Arabidopsis and the characterization of genetic variation for resistance or susceptibility. In
spite of the importance of disease to crop productivity, almost nothing is known about the
molecular mechanisms which specifically mediate the plant response to pathogens. Thus, the
rapid progress towards isolation of several disease resistance loci in Arabidopsis promises to
have direct practical applications in agriculture.
Pseudomonas syringae, P. cichorii, and Xanthomonas campestris strains that develop
compatible or incompatible interactions in Arabidopsis have been identified. At least two cloned
bacterial avirulence (avr) genes, avrRpt2 and avrRpm1, elicit the hypersensitive necrosis
response in an ecotype specific manner in Arabidopsis. At least four mutant alleles of a nuclear
locus in Arabidopsis fail to give a hypersensitive response when infiltrated with P. syringae
carrying the avrRpt2 gene, and an allele of a nuclear locus, found as a naturally occurring
variation in Arabidopsis ecotypes from the Kranz collection, confers resistance to avrRpm1.
Efforts are now underway in several laboratories to clone these putative resistance genes using
chromosome walking techniques. At a recent meeting in Köln (Germany) several laboratories
reported that they had identified RFLP markers within 1 to 2 cM of disease resistance genes and
the laboratory of J. Dangl (Köln, Germany) reported the isolation of a YAC fragment which
appears to carry the resistance gene RPM1.
Substantial progress has also been made in defining genes which confer susceptibility or
resistance to two obligate fungal biotrophs Albugo candida and Peronospora parasitica.
Screening of Arabidopsis accessions from the public strain collections identified lines which
were resistant or susceptible to Peronospora, Albugo and the powdery mildew pathogen
Erysiphe cruciferum. A hypothetical model involving eight resistance loci has been proposed to
explain differential interactions between six accessions and six isolates of Peronospora. One of
the loci has been mapped near gl1 on chromosome 3. Two other loci, RPP2 and RPP4, have
been located near to each other on chromosome 4. In addition, thirteen Arabidopsis accessions
have been found on which sporulation by Albugo does not occur or is delayed. The first
resistance locus (RAC1) is currently being mapped.
A variety of viruses are infectious in Arabidopsis, including cauliflower mosaic virus and
turnip crinkle mosaic virus, and at least one virus-resistant ecotype has been identified.
Arabidopsis has also been shown to be a good host for several sedentary root nematodes
including the cyst-knot nematode Heterodera schachtii and the root-knot nematode Meloidogyne
incognita. One advantage of Arabidopsis for the study of nematode-root interactions is that the
roots are sufficiently transparent to observe the entire life cycle of the nematodes using video-
enhanced contrast light microscopy. This has provided a unique opportunity to observe the
infection cycle in intact living tissue for the first time.
In addition to the direct analysis of disease resistance mechanisms involving major genes,
the rapidly expanding genetic resources in Arabidopsis provide unique opportunities to examine
the role of variation in other aspects of plant biology. For instance, ethylene has been

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hypothesized to play a role both in disease resistance and in disease susceptibility. This was
tested using isogenic virulent and avirulent bacterial pathogens, and mutants of Arabidopsis
thaliana altered in ethylene physiology. Ethylene-insensitive ein1 and ein2 mutants of
Arabidopsis were resistant to P. syringae pv. tomato made avirulent by addition of the cloned
avirulence genes avrRpt2, avrRpm1, or avrB, suggesting that ethylene is not required for active
resistance against avirulent bacteria. As another example, the structure of an Arabidopsis
phytoalexin (camalexin) was determined and mutants deficient in the accumulation of camalexin
were isolated. The availability of mutants should facilitate a direct and unequivocal test of the
role of phytoalexins in disease resistance to a broad spectrum of bacterial and fungal pathogens.
In addition, mutants have been isolated that show accelerated cell death symptoms (acd mutants)
when infected with bacterial or fungal pathogens.

C. Genes Cloned and Sequenced

The most important method for the isolation of new genes in Arabidopsis has become the
use of T-DNA tagging developed by K. Feldmann and D. Marks. At present, there are
approximately 13,000 lines in the collections maintained by DuPont and the two Arabidopsis
Resource Centers, each of which has a randomly inserted T-DNA. Approximately 30% of the
mutations observed in these lines are due to the insertion of T-DNA into a gene which can then
be readily cloned. The relative ease with which T-DNA tagged genes can be cloned has led to
rapid progress in understanding several previously intractable problems. For instance, T-DNA
tagging and related methods have been exploited with spectacular success to isolate a number of
genes involved in floral morphogenesis. During the past year at least 3 new genes involved in
flower development (APETALA3, LEAFY, PISTILLATA) have been isolated and a large group
of new enhancers, suppressors, and potentially tagged new flower mutations have been isolated
by E. Meyerowitz and collaborators (Pasadena, CA). In parallel with the isolation of genes by
genetic methods, a relatively large number of genes of already known function have been
isolated and characterized. For instance, because of the rapidly expanding interest in
pathogenesis in Arabidopsis, a variety of defense-related genes have been isolated for use in
monitoring reactions to pathogens. These include PAL1 (phenylalanine ammonia lyase), CHS
(chalcone synthase), BGL1, BGL2, BGL3 (ß-1,3-glucanases), GST1 (glutathione-S-transferase),
SOD1 (superoxide dismutase), and LOX1 (lipoxygenase). Several calmodulin-like proteins,
chitinase and HMG CoA reductase genes have been cloned. The number of cloned and
sequenced genes in the nucleic acid databases currently totals almost 300.

D. Large Scale Sequencing

The EC (European Community) is planning to fund a major initiative on sequencing


Arabidopsis. The working title is ESSA - European Scientists Sequencing Arabidopsis. Some
funding will be allocated to small scale sequencing in which well-characterized regions of the
genome will be sequenced at a rate of 25 kb/year. It is expected that grant proposals to sequence
cDNA clones on a similar scale will also be solicited. A small number of grants will be awarded
to undertake medium-scale sequencing of contiguous regions of approximately 1,000,000 bp per
year. It is estimated that there are currently three European laboratories which possess the
infrastructure to mount medium scale sequencing efforts. A medium-scale sequencing project is

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also underway in the laboratory of H. Goodman (Boston, MA). In preliminary feasibility
studies, 35 kb of contiguous sequence has been completed.
Large-scale cDNA sequencing has recently begun in France and the USA. The French
cDNA sequencing initiative is supported by CNRS. This project, in which 8 groups are
participating, is part of the French Government's initiative on Genome Projects. Each group has
prepared its own cDNA library corresponding to the biological problem it is interested in, and
has started to sequence, from both ends, cDNA clones selected either at random or after
differential screening. Nine months after the start of this program, approximately 600 clones
have been partially sequenced. Although limited, the available sequence information has
allowed the identification of slightly more than 200 genes by comparison with sequences already
in databanks. About half of them had not been identified previously in plants. The initial goal is
to partially sequence 3-4,000 clones. The general idea is to use these tagged sequences to
establish a transcription map of the Arabidopsis genome and to facilitate the genome sequencing
projects by providing the corresponding cDNA clones. These efforts will be integrated with the
EC BIOTECH program mentioned above and, through the use of common database (AAtDB),
with the other programs in the world. The coordinator of the program is Bernard Lescure
(Laboratoire de Biologie Moléculaire des Intéractions Plantes-Microrganismes, INRA-CNRS
B.P. 27, 31326 Castanet-Tolosan).
In the U.S., a cDNA sequencing laboratory has also been established at Michigan State
University with support from the U.S. Department of Energy Biological Energy Research
Program and the State of Michigan. The sequencing project was organized as a joint project by
ten laboratories at MSU and is managed by T. Newman. Sequencing reactions are performed by
an ABI Catalyst robot and resolved on ABI 373A Sequenators. Sequencing has begun on
random clones from a cDNA library representing roots, shoots, leaves, and etiolated seedlings.
The MSU project anticipates achieving a throughput rate of 200 partial cDNA sequences
(approximately 500 bp each) per week by the end of the year. The sequences are being
deposited on a daily basis in dbEST, a new public access database for expressed sequence tag
(EST) sequences operated by the National Center for Biotechnology, National Library of
Medicine. This database utilizes preprocessing and filtering tools that make EST analysis more
efficient and effective than the tools available in the larger databases (eg. GenBank). For more
information about dbEST, contact Mark Boguski at boguski@ncbi.nlm.nih.gov (Internet) or
301-496-2475 (phone). In addition, sequences will be analyzed for the presence of known
motifs and homology to known proteins by E. Retzel and colleagues at the University of
Minnesota. Results of these analyses will be deposited in the AIMS database.

2. Technology Development

A. Transformation Methods

During the past year the number of laboratories in which transformation has become
routine has greatly expanded due to the identification of several races of Arabidopsis which
respond well to improved transformation protocols. In addition, there have been several reports
that a method of Agrobacterium-mediated transformation reported several years ago by Hong-
Gil Nam (Korea) has been implemented in several laboratories. This method, which is an
adaptation of the method used by Feldmann to generate T-DNA inserts, involves inoculating

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decapitated plants with A. tumefaciens. Approximately one plant in five reportedly produces at
least one transformed progeny from such an inoculation. The method may have the potential
advantage of being race-insensitive and requiring substantially less effort than tissue-culture
based methods.
Important progress has been made during the past year in methods for large-scale
transformation of protoplasts and regeneration of transformed plants. The development of these
capabilities represents tangible progress toward the much needed methods for homologous gene
replacement and, possibly, the development of PACs (plant artificial chromosomes).

B. New Gene Cloning Methods

a. Chromosome walking

One of the important advantages of Arabidopsis is the small genome size and the
relatively low abundance of interspersed repetitive DNA. This makes it feasible to undertake
gene isolation by chromosome walking from flanking RFLP markers. There are currently
approximately 40 chromosome walking efforts underway worldwide. Several of these have
recently resulted in the isolation of functional genes which have complemented the mutations
used to mark the genes. In one instance, V. Arondel (East Lansing, MI) and collaborators
cloned a gene encoding an intractable enzyme in lipid metabolism by mapping a mutation
affecting membrane lipid composition and using the flanking RFLPs to isolate YACs covering
the mutation. The identified YACs were used to screen a cDNA library, and the gene identified
by transforming cDNA clones into the mutant in order to identify a clone which complemented
the mutation. In another instance, M. Estelle and collaborators (Bloomington, IN) isolated a
gene involved in mediating the response to exogenous auxin by identifying YACs covering the
axr1 mutation, then using a DNA rearrangement induced by the mutation to identify the region
of YAC DNA containing the gene. The isolation of these genes is expected to be followed in the
near future by the isolation of genes involved in pathogenesis, phytohormone and phytochrome
responsiveness, and regulation of flowering among other things. At least four walks have
covered regions of the genome of between 1000 and 1300 kb. Thus, although most scientists
involved in the walking efforts have found the approach to be expensive and demanding in many
respects, the fact that it is possible at all represents important progress. It is no longer necessary
to abandon an interesting mutation because of the lack of a method for isolating the gene. If the
mutation is sufficiently interesting, it can be cloned.
Progress on several fronts promises to facilitate the cloning of genes by map-based
cloning methods. Because of the participation of a relatively large number of laboratories in
chromosome walking efforts, the genetic maps in the regions of the walks have been
significantly refined and the YACs covering these regions defined. This process has already
resulted in the identification of YACs covering more than one third of the genome. As this
process continues, it will be increasingly possible to identify the YACs covering a region of the
genome by reference to a database. In addition, the recent development by J. Ecker's laboratory
(Philadelphia, PA) of a YAC library with significantly increased insert size will greatly facilitate
the development of an overlapping YAC library of the entire genome.

b. Insertional mutagenesis

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The most efficient method of cloning genes has been the use of T-DNA tagging. Mutants
from a major collection of lines carrying T-DNA inserts, which is maintained by DuPont Central
Research (Wilmington, DE), was distributed to 135 laboratories around the world during the past
year. The pooled transformed lines and individual visible mutants, generated by Agrobacterium
seed transformation in the Feldmann laboratory, are being distributed through the Nottingham
and Ohio stock centers. A total of 5,260 transformants, including 460 transformants generated at
Zoecon in 1986-87, and 4,800 generated at the University of Arizona in 1991, are available at
present. The transformed families have been subjected to a preliminary screen. Most of the
mutant families identified in these preliminary screens have been or are being sent to the two
stock centers.
Approximately 800 putative mutants were identified in the preliminary screen. These
include:

>60 flower mutants


>150 embryo-defective mutants
~150 size variants
>50 seedling-lethals
>100 pigment mutants
>100 other various morphological mutants.

The mutants listed above are currently available from the stock center or will be in the next
month.
A great deal of work has been involved in creating an active transposon for Arabidopsis.
The first products of this effort have been realized by the isolation of several Ds-caused mutants
by the groups of C. Dean and G. Coupland (Norwich, U.K.). Also, in the process of
characterizing a mutation which arose in one of the Ac lines generated by the Dean group, a new
transposon was isolated in a spontaneous mutant of the chl1 locus. Work in the laboratory of N.
Crawford (La Jolla, CA) indicates that this transposon is apparently active and may be useful as
an endogenous tagging system.

3. Biological Resource Centers

One of the initial objectives of the genome project was realized this year with the
opening of the Arabidopsis Biological Resource Center at Ohio State University (ABRC), in
addition to the previously established Nottingham Arabidopsis Stock Center (NASC) at the
University of Nottingham, U.K., and the DNA Resource Center in Köln, Germany. The
Nottingham Center was established under the direction of B. Mulligan and M. Anderson with
five years of funding from both the U.K. Agricultural and Food Research Council and the EC
BRIDGE program. The OSU Center, which is directed by R. Scholl and K. Davis, was
established with a five year grant from the National Science Foundation. The Köln Center
directed by J. Dangl is supported by EC-BRIDGE program. NASC and ABRC are operated as
parallel collections which closely coordinate the maintenance and collecting of plant materials.
As a general rule, it is expected that ABRC will service North America and NASC will service
the rest of the world.
The primary functions of these resource centers are to acquire, preserve and distribute
seed and DNA stocks for use by the research community. ABRC also collects and disseminates

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information concerning Arabidopsis and will provide a fully computerized stock
information/ordering system. In this regard, the Center is cooperating with S. Pramanik of
Michigan State University to develop a comprehensive database for Arabidopsis, AIMS
(Arabidopsis Information Management System). The Center, in addition to the co-principal
investigators, employs four technical staff.
The two seed stock collections currently maintain the Koornneef mutant and marker line
collection, a collection of approximately 400 T-DNA insertion lines from C. Koncz (Köln,
Germany), a collection of 5,000 T-DNA tagged lines from K. Feldmann (Tucson, AZ), the
collection of ecotypes and mutants previously maintained by A. Kranz (Frankfurt-am-Main,
Germany), and many mutant and marker lines contributed by other laboratories. In addition, the
Centers distribute 100 recombinant inbred lines contributed by P. Scolnik (Wilmington, DE).
Another 300 recombinant inbred lines from another cross will be contributed by C. Dean
(Norwich, U.K.) in October, 1992. ABRC is working with G. Redei (Columbia, MO) to transfer
his collection to the Ohio Center.
During 1991, the Nottingham Center distributed 791 seed lines and during the first five
months of 1992, more than 3,000 packets of seed were distributed to various locations around
the world. Since the ABRC began accepting orders on April 20, 1992 more than 12,000 seed
stocks had been distributed by August 1, 1992. Major contributions to these numbers are
represented by the Du Pont recombinant inbred lines and the Feldmann Agrobacterium mutants
and transformant pools. In addition, large numbers of tester lines, individual mutants and
T-DNA insertion mutants have also been shipped. It is obvious that there exists a large demand
for these resources.
Catalogs of the collections are available from the resource centers upon request. In
addition, as noted above, the AIMS database currently under development will be utilized to
keep the ABRC stock information current, for direct ordering of stocks and as a general
information resource on Arabidopsis.
Requests for stocks from the stock centers can be submitted by any of several means:
Specifically, a completed order form can be mailed, faxed or e-mailed to the Center. The
addresses for ordering are:

Biological Resource Center at Ohio State, 1735 Neil Avenue, Columbus, OH 43210, USA fax:
614-292-0603
telephone 614-292-9371
e-mail, seeds: seeds@genesys.cps.msu.edu (for seed orders)
e-mail, DNA: dna@genesys.cps.msu.edu (for DNA orders)
(NOTE: For e-mail orders, type "STOCK ORDER" in the subject line.)
The ABRC Stock list is deposited in the directory, Public/BIOSCI/ARABIDOPSIS. The file
name is ABRC_stock_lists and it can be recovered by anonymous FTP from genbank.bio.net.

For the Nottingham Center, contact:

Dr. M. Anderson, NASC, Department of Life Science, University of Nottingham, University


Park, Nottingham, NG7 2RD, UK
Telephone: 44-602-791216
Fax: 44-602-513251
E-mail: PLZMLH@VAX.CCC.NOTTINGHAM.AC.UK

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In addition to seed stocks, ABRC also maintains samples of plasmid, cosmid and YAC
clones. These include the mapped RFLP phage (from E. Meyerowitz) and cosmids (H.
Goodman), the YAC libraries produced by E. Grill (E. Lansing, MI) and E. Ward (Research
Triangle, NC) and individual genes. The Center accepts deposits of any cloned and
characterized Arabidopsis gene. In the three months of operation, ABRC has distributed 517
stocks of cosmid and phage clones and nine complete YAC libraries have been distributed.
Anticipated DNA acquisitions include more RFLP clones, genomic and cDNA libraries, and
individual clones. The DNA Resource Center maintained by J. Dangl at the Max Planck-Institut
für Züchtungsforschung in Köln (Germany) also distributes cDNA and genomic libraries which
have been deposited by members of the community. The Köln Center filled more than 30
requests for RFLP probes, YAC libraries and phage libraries in 1991 and 16 in 1992, so far. For
information about the Köln collection contact J. Dangl at DANGL@VAX.MPIZ-
KOELN.MPG.DBP.DE (E-mail) or 49-221-5062-613 (Fax).

4. Informatics
Because of the explosion of information about Arabidopsis, it has become essential to
have mechanisms to collect and organize the mass of new information. A tangible step forward
was implemented with the recent release of the Arabidopsis genome database, AAtDB (See
Appendix I). AAtDB (An Arabidopsis thaliana Data Base) uses the excellent ACeDB software
written by Richard Durbin (MCR-LMB, U.K.) and Jean Thierry-Mieg (CNRS, France). AAtDB
is funded by the U. S. Department of Agriculture Plant Genome Research Program through the
National Agricultural Library and is maintained by H. Goodman and colleagues at the
Massachusetts General Hospital and Harvard University in Boston. AAtDB is available without
charge via Internet network transfer.
The ACeDB software allows the user to browse information by simply pointing and
clicking with the workstation mouse. A powerful query facility is also available.

Currently AAtDB contains:

- The Hauge/Goodman cosmid/YAC physical map including >14,000 cosmid clones.


- Genetic markers, both RFLP and classical markers.
- Unified Genetic Map, including both the Goodman and Meyerowitz RFLP markers and
classical genetic markers.
- Primary F2 mapping database from the Goodman and Meyerowitz RFLP mapping
projects.
- Primary two point recombination data from M. Koornneef.
- A strain catalog including all strains and clones available from the Nottingham Stock
Centre and the ABRC at Ohio State University.
- Bibliographic citations from 1964 to present, currently numbering over 2,700.
- List of Arabidopsis researchers including mail address, phone number, FAX number and
electronic mail address. Currently information on over 500 colleagues is included.
- Green Book. The Green Book by Meyerowitz and Pruitt has been updated and integrated
into many parts of the database, including phenotype and allele descriptions.
- All DNA sequences from GenBank, currently there are over 300 sequences.
- BLASTX defined peptide sequence homologies.

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- REBASE restriction enzyme database maintained by R. Roberts.
- Graphical displays of all Genetic Maps, Physical Maps, and DNA Sequence features and
homologies.
- Scanned images of RFLP autoradiograms and photographs of mutant phenotypes.
The database presents the information in separate windows. There are many paths to any
piece of information allowing the user to easily navigate the connections between the various
types of information. As information about the Arabidopsis genome is expanded, AAtDB will
be extended and enhanced via periodic updates.
The database currently requires a Unix workstation running X-Windows. Versions of the
ACeDB database software are available for Sun Microsystems SPARCstations, Digital
Equipment's DECstation, Silicon Graphics Iris series and NeXT workstations. A printed manual
"An Introduction to ACeDB: For AAtDB--A. thaliana database" is available on request from the
MGH group. A MacIntosh version of the ACeDB software is under development. A public
release version is expected within six months. For more information contact Mike Cherry or
Sam Cartinhour at FAX number 617-726-6893 or via electronic mail at
curator@frodo.mgh.harvard.edu. By mail, contact Sam Cartinhour, J. Michael Cherry or
Howard M. Goodman, Department of Molecular Biology, Massachusetts General Hospital and
Department of Genetics, Harvard Medical School, Boston, MA 02114, USA.
A second database which will be available on-line via the Internet has been developed by
a collaboration between the OSU Resource Center and the laboratory of S. Pramanik (E.
Lansing, MI). The database, which is written in the very powerful Sybase language, will be
directly accessible by FTP via the Internet from any computer platform (i.e., Mac, PC or
workstation). The first release, which is currently being tested at several sites, will contain the
information on stocks in the resource centers and will permit ordering of seeds or DNA via
electronic media (See Appendix II). The full system, which should be implemented around the
end of 1992, will have complete genetic, phenotypic and other important data for germplasm,
relevant information for all clones, consensus genetic and physical maps and mapping data, and
bibliographical information. The feature of this database that distinguishes it from other species
genome databases is the attempt to incorporate detailed, atomized phenotypic information and to
allow complex searching involving phenotype. The MGH and MSU groups collaborate and
coordinate their activities in order to serve efficiently the Arabidopsis research community.
In order that the cDNA sequences produced by any large-scale sequencing efforts be of
the broadest possible utility, it is essential that they be deposited in a public access database.
However, partial cDNA sequence data is qualitatively different from traditional types of entries
in the general purpose sequence libraries. For instance, because the coding sequence and reading
frame of a partial cDNA sequence is not known for an EST (expressed sequence tag) in advance,
searching an EST collection for deduced amino acid sequence homology requires translating all
six frames with an amino acid query sequence, a function which is not routinely available on the
other databases. In addition, the map location of an EST, and not its relationship to known
sequences, may be the information of interest and this information is not explicitly retrievable
from the general purpose sequence collections. Because of the parallel need for such a database
structure in other genome projects, the National Center for Biotechnology Information (NCBI)
of the National Library of Medicine / the National Institutes of Health, has created a specialized
database called "dbEST" (data base of Expressed Sequence Tags). The Arabidopsis community
has been invited to deposit partial cDNA sequences in this database. The large-scale sequencing
effort underway at Michigan State University will also deposit all sequences in this database.

14
For more information contact Mark Boguski at:

National Center for Biotechnology Information, National Library of Medicine, Bldg, 38A, NIH,
8600 Rockville Pike, Bethesda, MD 20894, USA
Phone: 301-496-2475
Fax: 301-480-9241.

The EC BRIDGE project has established a committee to evaluate the needs and
conditions for operation of an European node to collect, analyze, disseminate and store genome
data. The members of the committee are M. Bevan (chair), J. Giraudat, D. Inze, M. Zabeau, and
J. Dangl. Following several meetings, including a joint EC-US workshop in January 1992, a
study report has been prepared which outlines the proposed structure of an EC Arabidopsis
database.

5. Communication

Since the inception of the electronic Arabidopsis newsgroup two years ago, most of the
laboratories working on Arabidopsis, and many scientists with a primary research interest in
other organisms, have subscribed to the newsgroup. The newsgroup is distributed worldwide
through both USENET news (under the name of bionet.genome.arabidopsis) and through e-mail.
There are currently 402 e-mail subscribers, 299 subscribed through the U.S. BIOSCI distribution
center at GenBank/IntelliGenetics and 103 subscribed through the BIOSCI center at SERC
Daresbury Laboratory in the U.K. While it is difficult to estimate precisely the number of
people who access the group via USENET news, a survey on another BIOSCI newsgroup about
a year ago indicated that almost 50% of the readership used USENET. This suggests that the
total newsgroup readership could be approximately 800 people at sites around the globe. There
have been 344 postings to the newsgroup during the
period 1 January 1992 through 27 July 1992 (209 days) for a usage rate of 1.65 messages per
day. This compares to a total of 264 messages posted during 1991 (0.94 per day over the period
3/26/91 to 12/31/92). The usage rate is up by 76% compared to last year. While overall 1992
statistics for BIOSCI remain to be compiled, in 1991 the Arabidopsis newsgroup was the seventh
most active group. The most active newsgroup, BIONEWS, averaged 3.78 messages per day in
1991 compared to 0.94 for Arabidopsis. The current year has seen a continuing growth in the
use of most BIOSCI forums as more biologists become familiar with electronic communications.
An archive of postings is maintained for anonymous FTP at genbank.bio.net in the directory
pub/BIOSCI/. A listing of the available files follows:

-rwxrwxr-x 1 kristoff 275510 Jan 2 1992 1991 (archive for 1991)


-rwxrwxr-x 1 kristoff 594778 Jul 27 13:46 1992 (archive for 1992)
-rwxrwxr-x 1 kristoff 51562 Jul 10 14:25 ABRC_stock_lists
-rwxrwxr-x 1 kristoff 70170 Apr 6 11:58 arab-gen.list (subscribers as of early 1992)
-rwxrwxr-x 1 kristoff 34932 Apr 6 11:58 arab-gen.list.pt1
-rwxrwxr-x 1 kristoff 35238 Feb 4 15:46 arab-gen.list.pt2

Postings are also indexed in the general "biosci.src" WAIS source on the computer
genbank.bio.net. WAIS software indexes all text in every BIOSCI newsgroup posting and

15
allows users on the Internet to search for any text string and then retrieve messages bearing the
specified text.
In order to subscribe to the newsgroup send a message requesting information on how to
subscribe to biosci@net.bio.net.
A useful and enjoyable source of information for those working on Arabidopsis has been
the quarterly newsletter sponsored by the British Arabidopsis group and edited by David
Flanders (Norwich, U.K.). Unfortunately, because of the limited funding available to publish
this lively and interesting newsletter, its circulation has been limited. However, it is expected
that a large-circulation newsletter, edited by several people worldwide, will appear in the
forthcoming year and will continue in the same vein as the British newsletter (which was
variously known as Arabadabadopis, Arabidian Notes, Thale and Cress, Arabido and several
other pseudonyms).

6. Workshops and Symposia

An important mechanism for stimulating communication among plant biologists has been
the convening of a number of specialized and general meetings focused on the use of
Arabidopsis. An international ARAPANET (Arabidopsis Pathogenesis Network) workshop on
pathogenesis was organized by Jeff Dangl in Köln, Germany. The workshop was attended by
approximately 50 people, 20 of whom were working with bacteria, fungi, viruses, and
nematodes that infect Arabidopsis.
A workshop to discuss genome research and database issues was held at Massachusetts
General Hospital (Boston, MA) in January 1992. Approximately 20 representatives of the
efforts underway in the U.S. and five European nations participated in a roundtable discussion
which was primarily focused on database issues associated with large-scale sequencing efforts.
A major result of the meeting was broad consensus that large-scale cDNA sequencing projects
must be organized in such a way that the community at large has rapid access to the information
via electronic databases. The broad consensus on this issue obviated discussion about the
problems created by the practice of seeking patents for partial cDNA sequences which was
initiated by the National Institutes of Health. There was unanimous agreement among the
participants of the Boston workshop that no attempt should be made to patent partial Arabidopsis
cDNA sequences.
The Fourth International Conference on Arabidopsis Research, which was attended by
approximately 450 participants, was held in Vienna in June 1990. The 5th International
conference is scheduled to be held at Ohio State University in 1993. The tentative dates are
August 19 to 22. The organizing committee is Keith Davis, Ken Feldmann, Randy Scholl, and
Roger Hangarter. For information, contact:

Roger P. Hangarter, Department of Plant Biology, Ohio State University, 1735 Neil Avenue,
Columbus, OH 43206, USA
E-mail: hangarter.1@osu.edu

II. Status of National and Transnational Research Projects

1. Australia and New Zealand

16
New Zealand now has at least two groups working with Arabidopsis. One group in the
Pastoral Research Institute at Palmerston North (formerly DSIR), which has a primary interest in
improving white clover, is using Arabidopsis to identify root specific promoters for expressing
resistance genes for various pests such as "grass grub". This illustrates how Arabidopsis
research may be integrated with problems in crop plants. Another group at the School of
Biological Sciences, University of Auckland, is trying to isolate genes for aluminum tolerance
by screening EMS populations. A similar project is also underway at CSIRO in Canberra.

2. European Community

An EC grant has been received to set up a Concerted Action Program on plant-nematode


interactions. The program has 18 participating institutes from 9 EC countries with a research
focus on Arabidopsis. The program will not fund research directly but can be used for general
meetings, frequent bilateral exchange visits, joint publications etc. In short, it is designed to
stimulate nematology groups to collaborate more through the use of a common model system.
As noted earlier, proposals have been called by the EC for grants to conduct large-scale
sequencing of the Arabidopsis genome.

3. France

During the last year, Arabidopsis research has attracted additional French scientists and
the initial effort made in the preceding year by the CNRS is beginning to bear fruits. Not only
has the CNRS continued its support of Arabidopsis research, but now, both INRA and the
Ministry of Research and Space are also funding research in this area. This Government effort
helps French groups to participate actively in the world-wide project, and promising results are
being produced. Various programs are now funded for the next two years and one can estimate
an input of 1 million U.S. dollars per year specifically supporting Arabidopsis projects, not
including regular grants and salaries of about 50 scientists and 50 technicians, students or
postdocs. The major effect of these programs has been to encourage the various groups to work
together and to integrate their research more closely with the international community,
especially that from the EC.
The major scientific goal of the leading groups in France remains to study genes and
their biological functions, including regulation of cell cycle, analysis of seed formation and
maturation, hormone and signal transduction, response to environmental stress and
gametogenesis. Two landmarks should be mentioned; the cloning by chromosome positioning
of the abi3 locus by Jerome Giraudat and the isolation by complementation cloning in yeast of a
gene for a potassium transporter by Herve Sentenac. For the other genes under study, most of
the work is focused on characterization of promoters and gene organization within each locus.
In addition, three major programs have been launched. The first is a cDNA sequencing
project mentioned earlier in this report. An important outcome of the program is that the French
researchers were able to equip their laboratories with computer and automatic sequencing
facilities, which should facilitate long-range mapping and sequencing. A second program,
supported primarily by the Ministry of Research and Space, is designed to develop transgenic
lines tagged with various insertions such as Ac, Tnt1 or various T-DNA constructs. This is part
of the genetic approach to developmental biology. So far, about 3,000 independent lines have

17
been generated by 4 groups, but their screening is still fairly limited. Finally a third project,
which is mostly concerned with stress physiology, is primarily supported by INRA.

4. Japan

Laboratories working with Arabidopsis have continued to increase in Japan, and now
number more than 12. The research topics include mutational analyses of flower organogenesis,
responses to physical and chemical stimuli, virus infection, hormonal regulation, transcriptional
regulation, and transformation, as well as isolation and characterization of genes involved in
heat-shock responses, desiccation, lipid biosynthesis, signaling pathways, and transcription.
These studies are funded primarily by the Ministry of Education, Science, and Culture, with
some grants from the Science and Technology Agency, the Ministry of Agriculture, Forestry and
Fisheries, and some from private Foundations.
The training course on the standard experimental techniques on Arabidopsis research was
held at the National Institute for Basic Biology at Okazaki on November 25 - December 1, 1991.
The course included lectures, demonstration, and practice on cultivation and genetic crosses,
transformation, and DNA/mRNA extraction and blotting analysis, as well as special lectures and
demonstration on transformation techniques by Dr. K. Feldmann of University of Arizona and
by Dr. H-G Nam of Korea, and of tissue printing technique by Dr. J. Varner of Washington
University. Sixteen attendants (graduate students and young researchers) were selected from
nearly 60 applicants from universities, national laboratories and private industry. Most of the
attendants had no prior experience in working with Arabidopsis. Following the training course,
the 2nd Workshop on Arabidopsis Studies was held on December 2 and 3, at the National
Institute of Basic Biology with more than 80 participants. Dr. B. Hauge of Massachusetts
General Hospital gave a talk on physical mapping at this workshop.
A book describing fundamental techniques on Arabidopsis cultivation, genetic analyses,
and transformation was published as part of the practical laboratory technique book series
written in Japanese. The training courses and meetings are planned for the coming year as well.

5. Russia

Because of a long history of using Arabidopsis as an experimental organism in Russia,


significant collections of mutants exist. Contact has been established between Russian groups
and the curators of the stock centers and a tentative agreement has been reached to transfer the
Russian collection to the NASC and ABRC.

6. United Kingdom

After the success of the 1989 three-year AFRC Plant Molecular Biology Program, part of
which funded 34 Arabidopsis project grants, a smaller program, PMB II, starts October, 1992.
This will consist of nearly 40 grants, over one-third of which are expected to be on Arabidopsis.

7. United States and Canada

A US-Canada steering committee was elected by an electronic vote system. The steering
committee consists of five members, three of whom serve on the Multinational Science Steering

18
Committee. The national committee will coordinate Arabidopsis research activities in the U.S.
and Canada by facilitating open communication among researchers and serving as an advisory
body to various funding agencies.
The U.S. Department of Agriculture, the Department of Energy, the National Institutes
of Health, and the National Science Foundation (NSF) continue to support Arabidopsis research.
The four agencies provided approximately $15 million during the period from October 1, 1990
to September 30, 1991 for Arabidopsis research. This includes new funding of $4.5 million
allocated by Congress to NSF for Arabidopsis genome research. Most of the funds supported
individual research programs covering virtually all plant biology research topics. In addition, a
significant amount was spent in support of postdoctoral research fellowships, growth facilities,
and the development of databases, technique/methods, and biological resources.

III. Analysis and Recommendations: New Goals for the Coming Year

The original Long-range Plan for the Multinational Coordinated Arabidopsis Genome
Research Project (publication #: NSF 90-80) identified a series of goals for the progress of the
multinational Arabidopsis genome project. An ongoing goal was to achieve saturation
mutagenesis so as to associate a phenotype with as many genes as possible, and to develop a
transposon tagging system to facilitate the cloning of genes by insertional mutagenesis. This
goal continues to be realized by the explosive growth of new mutations, by the creation and
distribution of large collections of tagged mutants, and by tangible progress toward identification
of a useful transposon. The short-term goals were to create satisfactory YAC libraries and to
link up the YACs into an ordered array. Subsequent recommendations in the first progress
report included integration of the existing linkage maps, and the definition of a strategy for the
collation of sequence data worldwide and for the mapping of sequenced regions to a unique
physical map. Progress has been made on both fronts. New YAC libraries with larger inserts
and fewer chimeric clones have been developed as technology has improved, and substantial
progress has been made toward linking up the YACs. Also, the genetic maps have been
consolidated and are being refined by the small-scale mapping efforts which are underway in
many laboratories.
Completion of the ordered YAC maps should remain a high priority. Although many
laboratories are currently utilizing YACs for chromosome walking, the consensus opinion is that
walking is not a sufficiently efficient way to isolate most genes. However, if an ordered set of
YACs were available, it would only be necessary to obtain high resolution mapping information
for a mutation of interest in order to identify the YAC containing the gene. That would reduce
the complexity of the problem to simply sorting through about 100 kb of DNA in order to
identify the gene of interest by its ability to complement a mutation. In addition, as the number
of cloned genes increases, the most facile way of placing cloned genes on the genetic map would
be to hybridize the genes to an ordered YAC library rather than to map RFLPs associated with
the gene.
At present, because of the economies of scale which can be achieved in constructing an
ordered map, the work has become focused in a small number of laboratories. In view of the
central importance of this project, it may be useful to consider expanding the number of
laboratories involved in this project. The most efficient way of doing this is probably to
coordinate the burgeoning cDNA sequencing projects with the effort to link up the YACs. That

19
is, in order to identify the function of most randomly sequenced cDNAs, it will be necessary to
obtain additional information about the clones such as genetic map location. The only feasible
strategy for accomplishing this is to hybridize the cDNA clones to YAC filters, or vice versa.
Whenever the cDNA clone hybridizes to two YAC clones in a region of overlap, information on
linkage of the YACs will result. Thus, by hybridizing several thousand cDNA clones to the
YAC libraries, it will be possible to link up the YACs. Therefore, we recommend that support be
allocated to large-scale mapping of partially sequenced cDNA clones.
A long term goal of the genome project was the complete sequencing of all cDNAs and
the mapping of these cDNAs on the genetic map. This aspect of the project has been initiated
with the establishment of one US laboratory dedicated to cDNA sequencing, and at least seven
European laboratories with this goal. In view of recent improvements in automated sequencing
instruments, it is apparent that with a modest investment, it is possible to obtain partial sequence
information on a majority of the estimated 20,000 mRNAs in Arabidopsis within the next
several years. In view of the fact that it has become possible to identify an increasing number of
deduced amino acid sequences by sequence identity or homology to polypeptides of known
function, this project promises to be very productive. As noted above, mapping of partially
sequenced cDNAs should be an integral part of this aspect of the project. We recommend that
the cDNA sequencing and mapping be assigned support with the caveat that suitable
mechanisms be put in place to ensure that the entire community has rapid and unrestricted access
to the results of all large-scale sequencing efforts. As noted below, additional cooperation must
be implemented to avoid wasteful duplication of effort in this enterprise.
Because of the importance of an accurate genetic map to all map based cloning efforts, it
is recommended that support be allocated to the establishment of a genetic map committee
which would be responsible for collecting all sequence information and integrating the data into
a consensus map which would be distributed via the databases and newsletters.
A result of the US-EC Sequence Database Workshop in January 1992 was the
recognition that there are a number of important issues pertaining to the organization of
international Arabidopsis nucleotide sequence databases that should be resolved in the
immediate future by subsequent meetings. In particular, a central database should be designated
and the design of data descriptors and nomenclature resolved before a large stream of data has
already been acquired. Issues pertaining to release of information from individual laboratories
must be resolved and some attempt to utilize the same biological materials as sources of DNA
should be agreed upon.
In the original goals of the project, a high priority was assigned to the creation of
biological resource centers. Two stock Centers and two DNA repositories are now operating and
the preliminary evidence indicates that they will serve an important role in ensuring rapid and
continuing access to the biological materials and information as they are developed. The
development of two databases is also an important step forward in providing rapid access to the
explosion of information. We strongly endorse continuing support for these resources. In the
area of communications, the Arabidopsis newsgroup has become an important mechanism for
discussion of issues and the sharing of technical information. The success of this important
medium must be attributed in large part to the responsive and efficient contribution of Dave
Kristofferson, the Genbank manager of the BIOSCI network. It is to be hoped that support for
the BIOSCI network will remain strong and unabated. The one remaining aspect of
communication which must be addressed in the near future is the creation of a newsletter. The
newsletter is expected to differ from the electronic newsgroup by providing a readily accessible

20
record of selected items from the electronic newsgroup, and by the inclusion of concise notes
about scientific observations which are of relevance primarily to the Arabidopsis community.
We anticipate that a newsletter will appear in the forthcoming year and encourage anyone who
would be interested in participating as an editor to contact a member of the Multinational
Science Steering Committee.
In addition to the progress described in this report, Arabidopsis research has undergone a
major transition during the past several years. In effect, the organism has become the most
widely utilized experimental organism in experimental plant biology. Thus, to present a full
account of Arabidopsis research has become very similar to summarizing progress in plant
biology. The implication in the present context is that it suggests that in order to avoid losing
sight of the goals of the genome project, it may be appropriate to begin focusing the project
more narrowly on the goals which are directly related to the Arabidopsis genome. In particular,
we believe it is time to begin allocating increased support for large-scale sequencing of both
cDNA and genomic clones.
In the area of human resource development, the ongoing goals which we continue to
endorse, were the support of multinational postdoctoral fellowships and short term exchanges
and short term courses.
An important aspect of research on Arabidopsis has been the spirit of cooperation and
openness which has characterized the field. The Arabidopsis genome project will generate many
unique research resources (nucleotide sequences, mapping data, clones, strains, databases and
software). Some mechanisms have now been established to facilitate free and unrestricted
access to these resources by all members of the international committee. It is to be hoped that all
participants in Arabidopsis research, regardless of their affiliation, will continue to make their
research results and accomplishments readily available. One mechanism which we strongly
endorse is the deposition of samples of new mutants, libraries and cloned genes in the resource
centers.
Progress in Arabidopsis research has occurred, in part, because Arabidopsis is integrated
into general genome and plant biology research activities. The Arabidopsis research community
has been quick to take advantage of any new developments in biological sciences, and at the
same time, information and knowledge gained through studies of Arabidopsis have been utilized
by the general biological research community. Major progress on many previously intractable
problems has been made using Arabidopsis as a model system and we anticipate that we will
soon begin to see the application of these advances to directed improvement of plants of
economic significance. It is, therefore, of great importance that mechanisms for interrelating the
information gained in Arabidopsis research and research on other plants be fostered. One
promising mechanism for facilitating this interplay will be the development of common
standards in the databases which are being developed for many crop species such as maize,
wheat, soybean and tomato. Another facilitating mechanism is to ensure that committees which
review grant proposals and related matters concerning research in crop species include members
of the Arabidopsis community who can provide information about the current developments in
Arabidopsis research. For similar reasons, membership on committees which deal primarily
with Arabidopsis research should continue to have representation by colleagues with primary
research interests which extend beyond Arabidopsis.
Finally, we recommend continued and expanded commitment to all of the goals set forth
for the Multinational Arabidopsis thaliana Genome Research Project. Funding agencies around
the world should continue to recognize the value of the truly international nature of the

21
Arabidopsis genome project, and allocate sufficient resources to help achieve the important
international goals.

22
Appendix I: AAtDB Example Display
This figure illustrates examples of the windows used by the ACeDB software developed by Drs.
Richard Durbin and Jean Thierry-Mieg to display the many types of research information
contained within the Arabidopsis thaliana database, AAtDB. Clockwise from upper left:
AAtDB main window listing the different topics included in the database, YAC and cosmid
physical map correlated to the genetic map, unified Arabidopsis genetic map of RFLP and
classical markers, strain descriptions and ordering information for both the ABRC at Ohio State
and the Nottingham Stock Center in the U.K., protein sequences with features, and finally,
extensive bibliographic citations. Version 1.1 of AAtDB was released to the public in early July,
1992, after three months of beta testing at Arabidopsis laboratories around the world. (Courtesy
of Dr. Michael Cherry.)

23
Appendix II: AIMS Sample Screen
AIMS (Arabidopsis Information Management System) contains detailed information on many
aspects of Arabidopsis. Included are data on DNA and seed stocks (see table at the top of
sample screen) and the facility for direct order placement. The graphic display of genetic maps,
etc. also is possible (see map of available chromosome 5 mutants at the bottom of sample
screen). (Courtesy of Dr. Sakti Pramanik.)

24
Appendix III:

Partial Listing of Recent Arabidopsis Publications

The following represents a partial listing of Arabidopsis research papers published from January
1991 to August 1992. This list was downloaded from "Reference Update" (Research
Information Systems, Inc., Camino Corporte Center, 2355 Camino Vida Roble, Carlsbad, CA)
and updated by Chris Somerville.

cDNA clones encoding Arabidopsis thaliana and Zea mays mitochondrial chaperonin HSP60 and gene expression
during seed germination and heat shock, Prasad,T.K., Stewart,C.R. Plant Mol. Biol., 18:873-885 (1992)

Characterization of rps17, rpl9 and rpl15: Three nucleus-encoded plastid ribosomal protein genes,
Thompson,M.D., Jacks,C.M. Lenvik,T.R., Gantt,J.S. Plant Mol. Biol., 18:931- 944 (1992)

Identification of an Arabidopsis DNA-binding protein with homology to nucleolin, Didier,D.K., Klee,H.J. Plant
Mol. Biol., 18:977-979 (1992)

An Arabidopsis thaliana cDNA clone encoding a 17.6 kDa class II heat shock protein, Bartling,D., Buelter,H.,
Liebeton,K. Weiler,E.W. Plant Mol. Biol., 18:1007-1008 (1992)

A new homeobox-leucine zipper gene from Arabidopsis thaliana, Mattsson,J., Soederman,E., Svenson,M.,
Borkird,C., Engstroem, P. Plant Mol. Biol., 18:1019-1022 (1992)

Ion channels in Arabidopsis plasma membrane. Transport characteristics and involvement in light-induced voltage
changes, Spalding, E.P., Slayman,C.L., Goldsmith,M.H.M., Gradmann,D., Bertl,A. Plant Physiol., 99:96-102
(1992)

A role for membrane lipid polyunsaturation in chloroplast biogenesis at low temperature, Hugly,S., Somerville,
C.R. Plant Physiol. 99:197-202 (1992)

Isolation and characterization of a mutant of Arabidopsis thaliana resistant to Alpha-methyltryptophan,


Kreps,J.A., Town,C.D. Plant Physiol., 99:269-275 (1992)

An Arabidopsis thaliana gene with sequence similarity to the S-locus receptor kinase of Brassica oleracea.
Sequence and expression, Tobias,C.M., Howlett,B., Nasrallah,J.B. Plant Physiol., 99:284-290 (1992)

Exon-intron organization of the Arabidopsis thaliana protein kinase genes CDC2a and CDC2b, Imajuku,Y.,
Hirayama,T., Endoh, H., Oka,A. FEBS Lett., 304:73-77 (1992)

Genes encoding a histone H3.3-like variant in Arabidopsis contain intervening sequences, Chaubet,N.,
Clement,B., Gigot,C. J. Mol. Biol., 225:569-574 (1992)

Identification of two tungstate-sensitive molybdenum cofactor mutants, chl2 and chl7, of Arabidopsis thaliana,
LaBrie,S. T., Wilkinson,J.Q., Tsay,Y.-F., Feldmann,K.A., Crawford,N.M. Molec. Gen. Genet., 233:169-176
(1992)
Expression of variant nuclear Arabidopsis tRNASer genes and pre-tRNA maturation differ in HeLa, yeast and
wheat germ extracts, Beier,D., Beier,H. Molec. Gen. Genet., 233:201-208 (1992)

25
Phytoalexin accumulation in Arabidopsis thaliana during the hypersensitive reaction to Pseudomonas syringae pv
syringae, Tsuji,J., Jackson,E.P., Gage,D.A., Hammerschmidt,R., Somerville, S.C. Plant Physiol., 98:1304-1309
(1992)

Role of abscisic acid in the induction of desiccation tolerance in developing seeds of Arabidopsis thaliana,
Meurs,C., Basra, A.S., Karssen,C.M., Van Loon,L.C. Plant Physiol., 98:1484-1493 (1992)

H-protein of the glycine decarboxylase multienzyme complex. Complementary DNA encoding the protein from
Arabidopsis thaliana, Srinivasan,R., Oliver,D.J. Plant Physiol., 98:1518-1519 (1992)

Isolation and characterization of a gene encoding a carboxypeptidase Y-like protein from Arabidopsis thaliana,
Bradley,D. Plant Physiol., 98:1526-1529 (1992)

Complementary DNA sequence of a low temperature-induced Brassica napus gene with homology to the
Arabidopsis thaliana kin1 gene, Orr,W., Iu,B., White,T.C., Robert,L.S., Singh,J. Plant Physiol., 98:1532-1534
(1992)

LEAFY controls floral meristem identity in Arabidopsis, Weigel, D., Alvarez,J., Smyth,D.R., Yanofsky,M.F.,
Meyerowitz,E.M.: Cell, 69:843-859 (1992)

A genetic model for light-regulated seedling development in Arabidopsis, Chory,J. Development, 115:337-354
(1992)

Germinal and somatic activity of the maize element Activator (Ac) in arabidopsis, Keller,J., Lim,E.,
James,D.W.,Jr., Dooner, H.K. Genetics, 131:449-459 (1992)

Strategies for mutagenesis and gene cloning using transposon tagging and T-DNA insertional mutagenesis,
Walbot,V. Ann. Rev. Plant. Physiol. Plant Mol. Biol., 43: 49-82 (1992)

Fusion events during floral morphogenesis, Verbeke,J.A. Ann. Rev. Plant. Physiol. Plant Mol. Biol., 43:583-598
(1992)

Mapping and sequencing of an actively transcribed Euglena gracilis chloroplast gene (ccsA) homologous to the
Arabidopsis thaliana nuclear gene cs(ch-42), Orsat,B., Monfort,A., Chatellard,P. Stutz,E. FEBS Lett.,
303:181-184 (1992)

A 126 bp fragment of a plant histone gene promoter confers preferential expression in meristems of transgenic
Arabidopsis, Atanassova,R., Chaubet,N., Gigot,C. Plant J., 2:291- 300 (1992)

Complementation of Saccharomyces cerevisiae auxotrophic mutants by Arabidopsis thaliana cDNAs, Minet,M.,


Dufour,M.-E., Lacroute, F. Plant J., 2:417-422 (1992)

Light-induced phosphorylation of a membrane protein plays an early role in signal transduction for phototropism
in Arabidopsis thaliana, Reymond,P., Short,T.W., Briggs,W.R., Poff,K.L.: Proc. Natl. Acad. Sci. USA

Genes that regulate plant development, Aeschbacher,R.A., Benfey, P.N. Plant Sci., 83:115-126 (1992)

Functional expression of a plant plasma membrane transporter in Xenopus oocytes, Boorer,K.J., Forde,B.G.,
Leigh,R.A., Miller, A.J. FEBS Lett., 302:166-168 (1992)

Identification of a single-copy gene encoding a Type I chlorophyll a/b-binding polypeptide of photosystem I in


Arabidopsis thaliana, Jensen,P.E., Kristensen,M., Hoff,T., Lehmbeck,J., Stummann, B.M., Henningsen,K.W.

26
Physiol. Plant., 84:561-567 (1992)

Analysis of multiple photoreceptor pigments for phototropism in a mutant of Arabidopsis thaliana, Konjevic,R.,
Khurana, J.P., Poff,K.L. Photochem. Photobiol., 55:789- 792 (1992)

A complete cDNA for adenine phosphoribosyltransferase from Arabidopsis thaliana, Moffatt,B.A.,


McWhinnie,E.A., Burkhart, W.E., Pasternak,J.J., Rothstein,S.J. Plant Mol. Biol., 18:653-662 (1992)

Cloning and sequencing of a cDNA encoding ascorbate peroxidase from Arabidopsis thaliana, Kubo,A., Saji,H.,
Tanaka,K., Tanaka, K., Kondo,N. Plant Mol. Biol., 18:691-701 (1992)

Nucleotide sequence of a cDNA encoding a protein kinase homologue in Arabidopsis thaliana, Mizoguchi,T.,
Hayashida,N., Yamaguchi- Shinozaki,K., Harada,H., Shinozaki,K. Plant Mol. Biol., 18:809-812 (1992)

Functional expression of a probable Arabidopsis thaliana potassium channel in Saccharomyces cerevisiae,


Anderson,J.A., Huprikar, S.S., Kochian,L.V., Lucas,W.J., Gaber,R.F. Proc. Natl. Acad. Sci.USA,

HD-Zip proteins Members of an Arabidopsis homeodomain protein superfamily, Schena,M., Davis,R.W. Proc.
Natl. Acad. Sci.USA

Embryonic mutants of Arabidopsis thaliana, Meinke,D.W. Dev. Genet., 12:382-392 (1991)

Improved method for the transformation of Arabidopsis thaliana with chimeric dihydrofolate reductase constructs
which confer methotrexate resistance, Kemper,E., Grevelding,C., Schell, J., Masterson,R. Plant Cell Reports,
11:118-121 (1992)

Effects of ionizing radiation on a plant genome: Analysis of two arabidopsis transparent testa mutations,
Shirley,B.W., Hanley,S., Goodman,H.M. Plant Cell, 4:333-347 (1992)

Hormone-resistant mutants of Arabidopsis have an attenuated response to Agrobacterium strains, Lincoln,C.,


Turner,J., Estelle,M. Plant Physiol., 98:979-983 (1992)

Shaking Arabidopsis thaliana, Sussman,M.R. Science, 256(5057): 619-619 (1992)

Production of polyhydroxybutyrate, a biodegradable thermoplastic in higher plants, Poirier,Y.P., Dennis,D.E.,


Klomparens,K., Somerville,C.R. Science 256:520-523 (1992)

Arabidopsis as a model system for studying plant disease resistance mechanisms, Innes,R., Bent,A., Whalen,M.,
Staskawicz,B. Ann. N.Y. Acad. Sci., 646:228-230 (1991)

Characterization of ADPglucose pyrophosphorylase from a starch- deficient mutant of Arabidopsis thaliana (L.),
Li,L., Preiss, J. Carboh. Res., 227:227-239 (1992)

Heterodimerization between light-regulated and ubiquitously expressed Arabidopsis GBF bZIP proteins,
Schindler,U., Menkens, A.E., Beckmann,H., Ecker,J.R., Cashmore,A.R. EMBO J., 11:1261-1273 (1992)

DNA binding site preferences and transcriptional activation properties of the Arabidopsis transcription factor
GBF1, Schindler, U., Terzaghi,W., Beckmann,H., Kadesch,T., Cashmore,A.R. EMBO J., 11:1275-1289 (1992)

Complementation of the cs dis2-11 cell cycle mutant of Schizosaccharomyces pombe by a protein phosphatase
from Arabidopsis thaliana, Nitschke,K., Fleig,U., Schell,J., Palme,K. EMBO J., 11:1327-1333 (1992)

Cloning and expression of an Arabidopsis nitrilase which can convert indole-3-acetonitrile to the plant hormone,

27
indole- 3-acetic acid, Bartling,D., Seedorf,M., Mithoefer,A., Weiler, E.W. Eur. J. Biochem., 205:417-424 (1992)

Leucine aminopeptidase from Arabidopsis thaliana--Molecular evidence for a phylogenetically conserved enzyme
of protein turnover in higher plants, Bartling,D., Weiler,E.W. Eur. J. Biochem., 205:425-431 (1992)

Multiple resistance to sulfonylureas and imidazolinones conferred by an acetohydroxyacid synthase gene with
separate mutations for selective resistance, Hattori,J., Rutledge,R., Labbe,H. Brown,D., Sunohara,G., Miki,B.
Molec. Gen. Genet., 232:167-173 (1992)

Behaviour of the maize transposable element Ac in Arabidopsis thaliana, Dean,C., Sjodin,C., Page,T., Jones,J.,
Lister,C. Plant J., 2:69-81 (1992)

Genetic and phenotypic characterization of cop 1 mutants of Arabidopsis thaliana, Deng,X.-W., Quail,P.H. Plant
J., 2:83-95 (1992)

terminal flower A gene affecting inflorescence development in Arabidopsis thaliana, Alvarez,J., Guli,C.L.,
Yu,X.-H., Smyth,D.R. Plant J., 2:103-116 (1992)

Regulated expression of the calmodulin-related TCH genes in cultured Arabidopsis cells: Induction by calcium
and heat shock, Braam,J. Proc. Natl. Acad. Sci.USA

Genes regulating the plant cell cycle: Isolation of a mitotic- like cyclin from Arabidopsis thaliana, Hemerly,A.,
Bergounioux, C., Van Montagu,M., Inze,D., Ferreira,P. Proc. Natl. Acad. Sci.USA,

A cold-regulated Arabidopsis gene encodes a polypeptide having potent cryoprotective activity, Lin,C.,
Thomashow,M.F. Biochem. Biophys. Res. Commun., 183:1103-1108 (1992)

Cotyledon cell development in Arabidopsis thaliana during reserve deposition, Mansfield,S.G., Briarty,L.G. Can.
J. Bot., 70:151-164 (1992)

SUPERMAN, a regulator of floral homeotic genes in Arabidopsis, Bowman,J.L., Sakai,H., Jack,T., Weigel,D.,
Mayer,U., Meyerowitz, E.M. Development, 114:599-615 (1992)

Sequence and organization of 5S ribosomal RNA-encoding genes of Arabidopsis thaliana, Campell,B.R.,


Song,Y., Posch,T.E. Cullis,C.A., Town,C.D. Gene, 112:225-228 (1992)

Cloning the Arabidopsis GA1 locus by genomic subtraction, Sun,T., Goodman,H.M., Ausubel,F.M. Plant Cell,
4:119-128 (1992)

Molecular analysis of an auxin binding protein gene located on chromosome 4 of Arabidopsis, Palme,K.,
Hesse,T., Campos, N., Garbers,C., Yanofsky,M.F., Schell,J. Plant Cell, 4: 193-201 (1992)

Mutations at the Arabidopsis chm locus promote rearrangements of the mitochondrial genome, Martinez-
Zapater,J.M., Gil-Vinuelas,P., Capel,J., Somerville,C.R. Plant Cell 4:889-899 (1992)
Structure of an mdr-like gene from Arabidopsis thaliana. Evolutionary implications, Dudler,R., Hertig,C. J. Biol.
Chem., 267(9):5882-5888 (1992)

A cDNA clone encoding HBP-1b homologue in Arabidopsis thaliana, Kawata,T., Imada,T., Shiraishi,H.,
Okada,K., Shimura,Y., Iwabuchi, M. Nucl. Acids Res., 20:1141-1141 (1992)

Isolation and characterization of genes encoding chaperonin 60Beta from Arabidopsis thaliana, Zabaleta,E.,
Oropeza,A., Jimenez,B., Salerno,G., Crespi,M., Herrera-Estrella,L. Gene, 111:175-181 (1992)

28
Enhancement of NaCl tolerance in Arabidopsis thaliana by exogenous L-asparagine and D-asparagine,
Lehle,F.R., Chen,F., Wendt, K.R. Physiol. Plant., 84:223-228 (1992)

Cold acclimation and cold-regulated gene expression in ABA mutants of Arabidopsis thaliana, Gilmour,S.J.,
Thomashow,M.F.: Plant Molec. Biol., 17:1233-1240 (1991)

cDNA sequence analysis and expression of two cold-regulated genes of Arabidopsis thaliana, Gilmour,S.J.,
Artus,N.N., Thomashow, M.F. Plant Mol. Biol., 18:13-21 (1992)

cDNA and derived amino acid sequence of a cytosolic Cu,Zn superoxide dismutase from Arabidopsis thaliana
(L.) Heyhn, Hindges,R. Slusarenko,A. Plant Mol. Biol., 18:123-125 (1992)

Sucrose synthase of Arabidopsis Genomic cloning and sequence characterization, Chopra,S., Del-favero,J.,
Dolferus,R., Jacobs, M. Plant Mol. Biol., 18:131-134 (1992)

Isolation and transcriptional competence of three tRNATrp genes from Arabidopsis thaliana L., Lin,T.-Y.,
March,R., Scanlon, S.R., Folk,W.R. Plant Mol. Biol., 18:159-160 (1992)

The glucosinolate-degrading enzyme myrosinase in Brassicaceae is encoded by a gene family, Xue,J.,


Lenman,M., Falk,A., Rask, L. Plant Mol. Biol., 18:387-398 (1992)

Molecular cloning approach for a putative ethylene receptor gene in Arabidopsis, Chang,C., Bleecker,A.B.,
Kwok,S.F., Meyerowitz, E.M. Biochem. Soc. Trans. 20:73-75 (1992)

Specific immune detection and partial purification of G-proteins from Arabidopsis thaliana, Clarkson,J.,
White,I.R., Millner, P.A. Biochem. Soc. Trans., 20:9S-9S (1992)

Crystallization and preliminary x-ray investigation of a ubiquitin carrier protein (E2) from Arabidopsis thaliana,
Cook,W.J., Jeffrey,L.C., Sullivan,M.L., Vierstra,R.D. J. Mol. Biol., 223:1183-1186 (1992)

Methylation sites in angiosperm genes, Gardiner-Garden,M., Sved,J.A., Frommer,M. J. Mol. Evol., 34: 219-230
(1992)

Significant CpG-rich regions in angiosperm genes, Gardiner- Garden,M., Frommer,M. J. Mol. Evol., 34: 231-245
(1992)

Structure and expression of AtS1, an Arabidopsis thaliana gene homologous to the S-locus related genes of
Brassica, Dwyer, K.G., Lalonde,B.A., Nasrallah,J.B., Nasrallah,M.E. Molec. Gen. Genet., 231:442-448 (1992)

Molecular cloning and sequence of cDNA encoding the pyrophosphate- energized vacuolar membrane proton
pump of Arabidopsis thaliana, Sarafian,V., Kim,Y., Poole,R.J., Rea,P.A. Proc. Natl. Acad. Sci.USA,

Sucrose mimics the light induction of Arabidopsis nitrate reductase gene transcription, Cheng,C.-L., Acedo,G.N.,
Cristinsin,M. Conkling,M.A. Proc. Natl. Acad. Sci.USA

The homeotic gene APETALA3 of arabidopsis thaliana encodes a MADS box and is expressed in petals and
stamens, Jack,T. Brockman,L.L., Meyerowitz,E.M. Cell, 68:683-697 (1992)

DNA binding activity of the arabidopsis G-box binding factor GBF1 is stimulated by phosphorylation by casein
kinase II from broccoli, Klimczak,L.J., Schindler,U., Cashmore,A.R. Plant Cell, 4:87-98 (1992)

Evidence for polyamine channels in protoplasts and vacuoles of Arabidopsis thaliana cells, Colombo,R.,
Cerana,R., Bagni, N. Biochem. Biophys. Res. Commun., 182: 1187-1192 (1992)

29
2-Trans-abscisic acid biosynthesis and metabolism of ABA-aldehyde and xanthoxin in wild type and the aba
mutant of Arabidopsis thaliana, Rock,C.D., Heath,T.G., Zeevaart,J.A.D. J. Exp. Bot., 43(247):249-256 (1992)

Global and local genome mapping in Arabidopsis thaliana by using recombinant inbred lines and random
amplified polymorphic DNAs, Reiter,R.S., Williams,J.G.K., Feldmann,K.A., Rafalski, J.A., Tingey,S.V.,
Scolnik,P.A. Proc. Natl. Acad. Sci.USA

Distribution of the rDNA and three classes of highly repetitive DNA in the chromatin of interphase nuclei of
Arabidopsis thaliana, Bauwens,S., Van Oostveldt,P., Engler,G., Van Montagu,M. Chromosoma, 101:41-48 (1991)

Linkage relationships of mutations that affect fatty acid composition in Arabidopsis, Hugly,S., Kunst,L.,
Somerville,C. J. Hered., 82:484-488 (1991)

Gene targeting in Arabidopsis thaliana, Halfter,U., Morris, P.-C., Willmitzer,L. Molec. Gen. Genet., 231:
186-193 (1992)

Expression of constitutive and tissue-specific acyl carrier protein isoforms in Arabidopsis, Hlousek-Radojcic,A.,
Post- Beittenmiller,D., Ohlrogge,J.B. Plant Physiol., 98:206- 214 (1992)

Occurrence of temperature-sensitive phenotypic plasticity in chlorophyll-deficient mutants of Arabidopsis


thaliana, Markwell, J., Osterman,J.C. Plant Physiol., 98:392-394 (1992)

Arabidopsis thaliana H1 histones--Analysis of two members of a small gene family, Gantt,J.S., Lenvik,T.R. Eur.
J. Biochem., 202:1029-1039 (1991)

Arabidopsis mutants deficient in polyunsaturated fatty acid synthesis. Biochemical and genetic characterization of
a plant oleoyl-phosphatidylcholine desaturase, Miquel,M., Browse,J. J. Biol. Chem., 267:1502-1509 (1992)

Genetic transformation of Arabidopsis thaliana zygotic embryos and identification of critical parameters
influencing transformation efficiency, Sangwan,R.S., Bourgeois,Y., Sangwan-Norreel,B. S. Molec. Gen. Genet.,
230:475-485 (1991)

Phytochrome B is not detectable in the hy3 mutant of Arabidopsis, which is deficient in responding to end-of-day
far-red light treatments, Nagatani,A., Chory,J., Furuya,M. Plant Cell Physiol., 32:1119-1122 (1991)
The hy3 long hypocotyl mutant of Arabidopsis is deficient in phytochrome B, Somers,D.E., Sharrock,R.A.,
Tepperman,J.M., Quail,P.H. Plant Cell, 3(12):1263-1274 (1991)

Overexpression of phytochrome B induces a short hypocotyl phenotype in transgenic Arabidopsis, Wagner,D.,


Tepperman,J.M., Quail, P.H. Plant Cell, 3(12):1275-1288 (1991)

Isolation and analysis of cDNAs encoding small GTP-binding proteins of Arabidopsis thaliana, Anai,T.,
Hasegawa,K., Watanabe, Y., Uchimiya,H., Ishizaki,R., Matsui,M. Gene, 108:259- 264 (1991)

Retention of phytochrome-mediated shade avoidance responses in phytochrome-deficient mutants of Arabidopsis,


cucumber and tomato, Whitelam,G.C., Smith,H. J. Plant Physiol., 139:119-125 (1991)

Isolation of mutants of Arabidopsis thaliana in which accumulation of tobacco mosaic virus coat protein is
reduced to low levels, Ishikawa,M., Obata,F., Kumagai,T., Ohno,T. Molec. Gen. Genet., 230(1-2):33-38 (1991)

The glutamine synthetase gene family of Arabidopsis thaliana: Light-regulation and differential expression in
leaves, roots and seeds, Peterman,T.K., Goodman,H.M. Molec. Gen. Genet., 230(1-2):145-154 (1991)

30
Insertional mutagenesis in Arabidopsis thaliana, Van Lijsebettens, M., Den Boer,B., Hernalsteens,J.-P., Van
Montagu,M. Plant Sci., 80(1-2):27-37 (1991)

Qualitative and quantitative genetic studies of Arabidopsis thaliana, Griffing,B., Scholl,R.L. Genetics,
129:605-609 (1991)

Cloning of a 16-kDa ubiquitin carrier protein from wheat and Arabidopsis thaliana. Identification of functional
domains by in vitro mutagenesis, Sullivan,M.L., Vierstra,R.D. J. Biol. Chem., 266(35):23878-23885 (1991)

Short-term competition for phosphate between two genotypes of Arabidopsis thaliana (L.) Heynh., Krannitz,P.G.,
Aarssen, L.W., Lefebvre,D.D. New Phytol., 119:389-396 (1991)

Sugar-dependent expression of the CHS-A gene for chalcone synthase from petunia in transgenic Arabidopsis,
Tsukaya,H., Ohshima, T., Naito,S., Chino,M., Komeda,Y. Plant Physiol., 97: 1414-1421 (1991)

Kinetics for phototropic curvature by etiolated seedlings of Arabidopsis thaliana, Orbovic,V., Poff,K.L. Plant
Physiol., 97:1470-1475 (1991)

Identification and properties of the major ribonucleases of Arabidopsis thaliana, Yen,Y., Green,P.J. Plant
Physiol., 97:1487-1493 (1991)

Electrogenic transport properties of growing Arabidopsis root hairs. The plasma membrane proton pump and
potassium channels, Lew,R.R. Plant Physiol., 97:1527-1534 (1991)

Nucleotide sequence of a cDNA clone encoding a Beta-amylase from Arabidopsis thaliana, Monroe,J.D.,
Salminen,M.D., Preiss, J. Plant Physiol., 97:1599-1601 (1991)

Phytochrome-deficient hy1 and hy2 long hypocotyl mutants of arabidopsis are defective in phytochrome
chromophore biosynthesis, Parks,B.M., Quail,P.H. Plant Cell, 3(11):1177-1186 (1991)

The FLO10 gene product regulates the expression domain of homeotic genes AP3 and PI in Arabidopsis flowers,
Schultz,E.A., Pickett, F.B., Haughn,G.W. Plant Cell, 3(11):1221-1237 (1991)

Repression of plant tissue culture growth by light is caused by photochemical change in the culture medium,
Hangarter,R. P., Stasinopoulos,T.C. Plant Sci., 79:253-257 (1991)

Phytochrome A overexpression inhibits hypocotyl elongation in transgenic Arabidopsis, Boylan,M.T., Quail,P.H.


Proc. Natl. Acad. Sci.USA

Correlation of apoproteins with the genes of the major chlorophyll a/b binding protein of photosystem II in
Arabidopsis thaliana: Confirmation for the presence of a third member of the LHC IIb gene family,
Morishige,D.T., Thornber,J.P. FEBS Lett., 293(1-2):183-187 (1991)

Isolation and characterization of hormone-autonomous tumours of Arabidopsis thaliana, Persinger,S.M.,


Town,C.D. J. Exp. Bot., 42(244):1363-1370 (1991)

Temporal and spatial development of the cells of the expanding first leaf of Arabidopsis thaliana (L.) Heynh.,
Pyke,K.A., Marrison,J.L., Leech,R.M. J. Exp. Bot., 42(244):1407-1416 (1991)

Nucleotide sequence and expression of a novel glycine-rich protein gene from Arabidopsis thaliana, Quigley,F.,
Villiot, M.-L., Mache,R. Plant Mol. Biol., 17:949-952 (1991)

Effect of chlorate treatment on nitrate reductase and nitrite reductase gene expression in Arabidopsis thaliana,

31
LaBrie, S.T., Wilkinson,J.Q., Crawford,N.M. Plant Physiol., 97: 873-879 (1991)

Molecular basis of imidazolinone herbicide resistance in Arabidopsis thaliana var Columbia, Sathasivan,K.,
Haughn,G.W., Murai,N. Plant Physiol., 97:1044-1050 (1991)

A mutant of Arabidopsis deficient in xylem loading of phosphate, Poirier,Y., Thoma,S., Somerville,C.,


Schiefelbein,J. Plant Physiol., 97:1087-1093 (1991)

Inward rectifying K+ channels in the plasma membrane of Arabidopsis thaliana, Colombo,R., Cerana,R. Plant
Physiol., 97: 1130-1135 (1991)

Physiology of hormone autonomous tissue lines derived from radiation-induced tumors of Arabidopsis thaliana,
Campell, B.R., Town,C.D. Plant Physiol., 97:1166-1173 (1991)

A myb gene required for leaf trichome differentiation in Arabidopsis is expressed in stipules, Oppenheimer,D.G.,
Herman,P.L., Sivakumaran, S., Esch,J., Marks,M.D. Cell, 67:483-493 (1991)

rDNA intergenic region from Arabidopsis thaliana. Structural analysis, intraspecific variation and functional
implications, Gruendler,P., Unfried,I., Pascher,K., Schweizer,D. J. Mol. Biol., 221:1209-1222 (1991)

Cloning and chromosomal mapping of nuclear genes encoding chloroplast and cytosolic
glyceraldehyde-3-phosphate-dehydrogenase from Arabidopsis thaliana, Shih,M.-C., Heinrich,P., Goodman,H.M.
Gene, 104:133-138 (1991)

Developmental and pathogen-induced activation of the Arabidopsis acidic chitinase promoter, Samac,D.A.,
Shah,D.M. Plant Cell, 3(10):1063-1072 (1991)

Identification of two cell-cycle-controlling cdc2 gene homologs in Arabidopsis thaliana, Hirayama,T.,


Imajuku,Y., Anai,T., Matsui,M., Oka,A. Gene, 105:159-165 (1991)

Characterization of the gene encoding the 10 kDa polypeptide of photosystem II from Arabidopsis thaliana,
Gil-Gomez,G., Marrero,P.F., Haro,D., Ayte,J., Hegardt,F.G. Plant Mol. Biol., 17:517-522 (1991)

A mutation in the arabidopsis TFL1 gene affects inflorescence meristem development, Shannon,S.,
Meeks-Wagner,D.R. Plant Cell, 3(9):877-892 (1991)

Overexpression of acetohydroxyacid synthase from Arabidopsis as an inducible fusion protein in Escherichia coli.
Production of polyclonal antibodies, and immunological characterization of the enzyme, Singh,B., Schmitt,G.,
Lillis,M., Hand,J.M., Misra,R. Plant Physiol., 97:657-662 (1991)

Effects of the gibberellin biosynthetic inhibitor uniconazol on mutants of Arabidopsis, Nambara,E., Akazawa,T.,
McCourt, P. Plant Physiol., 97:736-738 (1991)

A DNA transformation-competent Arabidopsis genomic library in Agrobacterium, Lazo,G.R., Stein,P.A.,


Ludwig,R.A. BioTechnology, 9(10):963-967 (1991)

Differential induction of 3-deoxy-D-arabino-heptulosonate 7- phosphate synthase genes in Arabidopsis thaliana


by wounding and pathogenic attack, Keith,B., Dong,X., Ausubel,F.M., Fink, G.R. Proc. Natl. Acad. Sci.USA

A genetic and physiological analysis of late flowering mutants in Arabidopsis thaliana, Koornneef,M.,
Hanhart,C.J., Van der Veen,J.H. Molec. Gen. Genet., 229:57-66 (1991)

Mutations affecting body organization in the Arabidopsis embryo, Mayer,U., Ruiz,R.A.T., Berleth,T., Misera,S.,

32
Juergens,G.: Nature, 353(6343):402-407 (1991)

Abscisic-acid-deficient mutants at the aba gene locus of Arabidopsis thaliana are impaired in the epoxidation of
zeaxanthin, Duckham, S.C., Linforth,R.S.T., Taylor,I.B. Plant Cell Environ., 14:601-606 (1991)

Biochemical genetics of plant secondary metabolites in Arabidopsis thaliana. The glucosinolates, Haughn,G.W.,
Davin,L., Giblin, M., Underhill,E.W. Plant Physiol., 97:217-226 (1991)

Reversible inactivation of a transgene in Arabidopsis thaliana, Mittelsten Scheid,O., Paszkowski,J., Potrykus,I.


Molec. Gen. Genet., 228(1-2):104-112 (1991)

Nucleotide sequence of a cDNA clone encoding chloroplast phosphoribuloki ase from Arabidopsis thaliana,
Horsnell,P.R., Raines,C.A.: Plant Mol. Biol., 17:183-184 (1991)

Nucleotide sequence of a cDNA clone encoding chloroplast fructose- 1,6-bisphosphatase from Arabidopsis
thaliana, Horsnell,P.R. Raines,C.A. Plant Mol. Biol., 17:185-186 (1991)

Transient expression of Beta-glucuronidase in Arabidopsis thaliana leaves and roots and Brassica napus stems
using a pneumatic particle gun, Seki,M., Komeda,Y., Iida,A., Yamada,Y., Morikawa, H. Plant Mol. Biol.,
17:259-263 (1991)

Ethylene binding in wild type and mutant Arabidopsis thaliana (L.) Heynh, Sanders,I.O., Harpham,N.V.J.,
Raskin,I., Smith, A.R., Hall,M.A. Annals Bot., 68:97-103 (1991)

Expression of the Arabidopsis floral homeotic gene AGAMOUS is restricted to specific cell types late in flower
development, Bowman,J.L., Drews,G.N., Meyerowitz,E.M. Plant Cell, 3(8): 749-758 (1991)

LEAFY, a homeotic gene that regulates inflorescence development in Arabidopsis, Schultz,E.A., Haughn,G.W.
Plant Cell, 3(8): 771-781 (1991)

Effect of light on the NADPH-protochlorophyllide oxidoreductase of Arabidopsis thaliana, Benli,M., Schulz,R.,


Apel,K. Plant Mol. Biol., 16:615-625 (1991)
Expression of nuclear tRNATyr genes from Arabidopsis thaliana in HeLa cell and wheat germ extracts,
Stange,N., Beier,D., Beier,H. Plant Mol. Biol., 16:865-875 (1991)

Separate signal pathways regulate the expression of a low-temperature- induced gene in Arabidopsis thaliana (L.)
Heynh., Nordin,K. Heino,P., Palva,E.T. Plant Mol. Biol., 16:1061- 1071 (1991)

The complete nucleotide sequence of the intergenic spacer region of an rDNA operon from Brassica oleracea and
its comparison with other crucifers, Bennett,R.I., Smith,A.G. Plant Mol. Biol., 16:1095-1098 (1991)

The aba mutant of Arabidopsis thaliana is impaired in epoxy- carotenoid biosynthesis, Rock,C.D.,
Zeevaart,J.A.D. Proc. Natl. Acad. Sci.USA

Nucleotide sequences of the mitochondrial genes trnS(TGA) encoding tRNASerTGA in Oenothera berteriana and
Arabidopsis thaliana, Binder,S., Knoop,V., Brennicke,A. Gene, 102:245-247 (1991)

Mapping genes essential for embryo development in Arabidopsis thaliana, Patton,D.A., Franzmann,L.H.,
Meinke,D.W. Molec. Gen. Genet., 227:337-347 (1991)

Rapid image analysis screening procedure for identifying chloroplast number mutants in mesophyll cells of
Arabidopsis thaliana (L. ) Heynh., Pyke,K.A., Leech,R.M. Plant Physiol., 96: 1193-1195 (1991)

33
Primary structures of Arabidopsis calmodulin isoforms deduced from the sequences of cDNA clones, Ling,V.,
Perera,I., Zielinski, R.E. Plant Physiol., 96:1196-1202 (1991)

Arabinose kinase-deficient mutant of Arabidopsis thaliana, Dolezal,O., Cobbett,C.S. Plant Physiol., 96:1255-1260
(1991)

Isolation, characterization, and chromosomal location of a new cab gene from Arabidopsis thaliana, Zhang,H.,
Hanley,S. Goodman,H.M. Plant Physiol., 96:1387-1388 (1991)

The effect of ethylene on the growth and development of wild- type and mutant Arabidopsis thaliana (L.) Heynh,
Harpham,N. V.J., Berry,A.W., Knee,E.M., Roveda-Hoyos,G., Raskin,I., Sanders, I.O., Smith,A.R., Wood,C.K.,
Hall,M.A. Annals Bot., 68:55-61 (1991)

Genetic analysis of pattern formation in the Arabidopsis embryo, Juergens,G., Mayer,U., Torres Ruiz,R.A.,
Berleth,T., Misera, S. Development, 112, Suppl. 1:27-38 (1991)

A genetic and molecular model for flower development in Arabidopsis thaliana, Meyerowitz,E.M., Bowman,J.L.,
Brockman,L.L., Drews, G.N., Jack,T., Sieburth,L.E., Weigel,D. Development, 112, Suppl. 1:157-167 (1991)

Analysis of two linked genes coding for the acyl carrier protein (ACP) from Arabidopsis thaliana (columbia),
Lamppa,G., Jacks, C. Plant Mol. Biol., 16:469-474 (1991)

Requirement of the auxin polar transport system in early stages of Arabidopsis floral bud formation, Okada,K.,
Ueda,J., Komaki, M.K., Bell,C.J., Shimura,Y. Plant Cell, 3:677-684 (1991)

Arabidopsis mutants lacking blue light-dependent inhibition of hypocotyl elongation, Liscum,E., Hangarter,R.P.
Plant Cell, 3:685-694 (1991)

Genes with homology to fungal and S-gene RNases are expressed in Arabidopsis thaliana, Taylor,C.B.,
Green,P.J. Plant Physiol., 96:980-984 (1991)

Systemic endopolyploidy in Arabidopsis thaliana, Galbraith, D.W., Harkins,K.R., Knapp,S. Plant Physiol.,
96:985- 989 (1991)

Gravitropism and starch statoliths in an Arabidopsis mutant, Saether,N., Iversen,T.-H. Planta, 184:491-497 (1991)

What the papers say Molecular genetics of floral development in Arabidopsis thaliana, Pruitt,R.E. BioEssays,
13:347- 349 (1991)

cop1 A regulatory locus involved in light-controlled development and gene expression in Arabidopsis,
Deng,X.-W., Caspar,T., Quail,P.H. Genes Devel., 5:1172-1182 (1991)

The centromere region of Arabidopsis thaliana chromosome 1 contains telomere-similar sequences, Richards,E.J.,
Goodman, H.M., Ausubel,F.M. Nucl. Acids Res., 19(12):3351-3357
(1991)

In vitro desaturation of monogalactosyldiacylglycerol and phosphatidylch line molecular species by chloroplast


homogenates, Norman, H.A., Pillai,P., St.John,J.B. Phytochemistry, 30:2217- 2222 (1991)

Phytochrome control of the tms2 gene in transgenic Arabidopsis: A strategy for selecting mutants in the signal
transduction pathway, Karlin-Neumann,G.A., Brusslan,J.A., Tobin,E.M. Plant Cell, 3:573-582 (1991)

In vivo random Beta-glucuronidase gene fusions in Arabidopsis thaliana, Kertbundit,S., De Greve,H.,

34
Deboeck,F., Van Montagu, M., Hernalsteens,J.-P. Proc. Natl. Acad. Sci.USA

Negative regulation of the arabidopsis homeotic gene AGAMOUS by the APETALA2 product, Drews,G.N.,
Bowman,J.L., Meyerowitz, E.M. Cell, 65:991-1002 (1991)

A novel class of plant proteins containing a homeodomain with a closely linked leucine zipper motif, Ruberti,I.,
Sessa,G. Lucchetti,S., Morelli,G. EMBO J., 10:1787-1791 (1991)

Effects of exogenous polyamines and difluoromethylornithine on seed germination and root growth of Arabidopsis
thaliana, Mirza,J.I., Bagni,N. Plant Growth Reg., 10:163-168 (1991)

Correction for non-linear relationships between root size and short term Pi uptake in genotype comparisons,
Krannitz,P.G. Aarssen,L.W., Lefebvre,D.D. Plant Soil, 133:157-167 (1991)

Relationships between physiological and morphological attributes related to phosphate uptake in 25 genotypes of
Arabidopsis thaliana, Krannitz,P.G., Aarssen,L.W., Lefebvre,D.D. Plant Soil, 133:169-175 (1991)

Cloning and characterization of an inorganic pyrophosphatase gene from Arabidopsis thaliana, Kieber,J.J.,
Signer,E.R.: Plant Mol. Biol., 16:345-348 (1991)

The role of homeotic genes in flower development and evolution, Coen,E.S. Ann. Rev. Plant. Physiol. Plant Mol.
Biol., 42:241-279 (1991)

Glycerolipid synthesis Biochemistry and regulation, Browse, J., Somerville,C. Ann. Rev. Plant. Physiol. Plant
Mol. Biol., 42:467-506 (1991)

Genetic interactions among floral homeotic genes of Arabidopsis, Bowman,J.L., Smyth,D.R., Meyerowitz,E.M.
Development, 112: 1-20 (1991)

Tryptophan mutants in Arabidopsis The consequences of duplicated tryptophan synthase Beta genes, Last,R.L.,
Bissinger,P.H., Mahoney,D.J., Radwanski,E.R., Fink,G.R. Plant Cell, 3: 345-358 (1991)

Phenotypic and genetic analysis of det2, a new mutant that affects light-regulated seedling development in
Arabidopsis, Chory,J., Nagpal,P., Peto,C.A. Plant Cell, 3:445-459 (1991)

Identification of the Arabidopsis CHL3 gene as the nitrate reductase structural gene NIA2, Wilkinson,J.Q.,
Crawford,N. M. Plant Cell, 3:461-471 (1991)

The Arabidopsis functional homolog of the p34cdc2 protein kinase, Ferreira,P.C.G., Hemerly,A.S., Villarroel,R.,
Van Montagu,M. Inze,D. Plant Cell, 3:531-540 (1991)

Circadian control of cab gene transcription and mRNA accumulation in Arabidopsis, Millar,A.J., Kay,S.A. Plant
Cell, 3:541- 550 (1991)

The microtubular cytoskeleton during development of the zygote, proembryo and free-nuclear endosperm in
Arabidopsis thaliana (L.) Heynh, Webb,M.C., Gunning,B.E.S. Planta, 184:187- 195 (1991)

Differential expression of the two Arabidopsis nitrate reductase genes, Cheng,C.-L., Acedo,G.N., Dewdney,J.,
Goodman,H.M., Conkling,M.A. Plant Physiol., 96:275-279 (1991)

Isolation and characterization of a gene encoding a PR-1-like protein from Arabidopsis thaliana, Metzler,M.C.,
Cutt,J.R. Klessig,D.F. Plant Physiol., 96:346-348 (1991)

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Gap junction protein homologue from Arabidopsis thaliana Evidence for connexins in plants, Meiners,S., Xu,A.,
Schindler, M. Proc. Natl. Acad. Sci.USA

Early embryogenesis in Arabidopsis thaliana. I. The mature embryo sac, Mansfield,S.G., Briarty,L.G., Erni,S.
Can. J. Bot., 69:447-460 (1991)

Early embryogenesis in Arabidopsis thaliana. II. The developing embryo, Mansfield,S.G., Briarty,L.G. Can. J.
Bot., 69:461-476 (1991)

Transient occurrence of extrachromosomal DNA of an Arabidopsis thaliana transposon-like element, Tat1,


Peleman,J., Cottyn, B., Van Camp,W., Van Montagu,M., Inze,D. Proc. Natl. Acad. Sci. USA

The ubiquitin-encoding multigene family of flax, Linum usitatissimum, Agarwal,M.L., Cullis,C.A. Gene,
99:69-75 (1991)

Characterization and evolution of napin-encoding genes in radish and related crucifers, Raynal,M., Depigny,D.,
Grellet,F., Delseny,M. Gene, 99:77-86 (1991)

Two genes, atpC1 and atpC2, for the gamma subunit of Arabidopsis thaliana chloroplast ATP synthase,
Inohara,N., Iwamoto,A., Moriyama,Y., Shimomura,S., Maeda,M., Futai,M. J. Biol. Chem., 266:7333-7338 (1991)

A novel seed protein gene from Vicia faba is developmentally regulated in transgenic tobacco and Arabidopsis
plants, Baeumlein, H., Boerjan,W., Nagy,I., Bassuener,R., Van Montagu,M., Inze, D., Wobus,U. MGG. Molec.
Gen. Genet., 225: 459-467 (1991)

Nucleotide sequences of two genomic DNAs encoding peroxidase of Arabidopsis thaliana, Intapruk,C.,
Higashimura,N., Yamamoto, K., Okada,N., Shinmyo,A., Takano,M. Gene, 98:237-241 (1991)

Mutants of Arabidopsis with altered regulation of starch degradation, Caspar,T., Lin,T.-P., Kakefuda,G.,
Benbow,L., Preiss,J., Somerville, C. Plant Physiol., 95:1181-1188 (1991)

A superfamily of Arabidopsis thaliana retrotransposons, Konieczny, A., Voytas,D.F., Cummings,M.P.,


Ausubel,F.M. Genetics, 127: 801-809 (1991)

Nuclear tRNATyr genes are highly amplified at a single chromosomal site in the genome of Arabidopsis thaliana,
Beier,D., Stange, N., Gross,H.J., Beier,H. Molec. Gen. Genet., 225:72-80 (1991)

Cis and trans-acting elements involved in the activation of Arabidopsis thaliana A1 gene encoding the translation
elongation factor EF-1Alpha, Curie,C., Liboz,T., Bardet,C., Gander,E. Medale,C., Axelos,M., Lescure,B. Nucl.
Acids Res., 19:1305-1310 (1991)

Embryonic lethals and T-DNA insertional mutagenesis in Arabidopsis, Errampalli,D., Patton,D., Castle,L.,
Mickelson,L., Hansen,K. Schnall,J., Feldmann,K., Meinke,D. Plant Cell, 3:149-157 (1991)

A quantification of the significance of assimilatory starch for growth of Arabidopsis thaliana L. Heynh,
Schulze,W., Stitt, M., Schulze,E.-D., Ekkehard Neuhaus,H., Fichtner,K. Plant Physiol., 95:890-895 (1991)

Metabolism of benzyladenine is impaired in a mutant of Arabidopsis thaliana lacking adenine


phosphoribosyltransferase activity, Moffatt,B., Pethe,C., Laloue,M. Plant Physiol., 95:900- 908 (1991)

The effect of genetically based differences in seed size on seedling survival in Arabidopsis thaliana
(Brassicaceae), Krannitz,P.G., Aarssen,L.W., Dow,J.M. Am. J. Bot., 78:446-450 (1991)

36
AGL1-AGL6, an Arabidopsis gene family with similarity to floral homeotic and transcription factor genes, Ma,H.,
Yanofsky,M. F., Meyerowitz,E.M. Genes Devel., 5:484-495 (1991)

Identification of Pseudomonas syringae pathogens of Arabidopsis and a bacterial locus determining avirulence on
both Arabidopsis and soybean, Whalen,M.C., Innes,R.W., Bent,A.F., Staskawicz, B.J. Plant Cell, 3:49-59(1991)
Induction of Arabidopsis defense genes by virulent and avirulent Pseudomonas syringae strains and by a cloned
avirulence gene, Dong,X., Mindrinos,M., Davis,K.R., Ausubel,F.M. Plant Cell, 3:61-72 (1991)

Chloroplasts of Arabidopsis thaliana homozygous for the ch- 1 locus lack chlorophyll b, lack stable LHCPII and
have stacked thylakoids, Murray,D.L., Kohorn,B.D. Plant Mol. Biol., 16:71-79 (1991)

T-DNA integration A mode of illegitimate recombination in plants, Mayerhofer,R., Koncz-Kalman,Z.,


Nawrath,C., Bakkeren, G., Crameri,A., Angelis,K., Redei,G.P., Schell,J., Hohn,B., Koncz,C. EMBO J.,
10:697-704 (1991)

Structure and molecular evolutionary analysis of a plant cytochrome c gene Surprising implications for
Arabidopsis thaliana, Kemmerer,E.C., Lei,M., Wu,R. J. Molec. Evolution, 32:227-237 (1991)

Purification and characterization of an antifungal chitinase from Arabidopsis thaliana, Verburg,J.G., Huynh,Q.K.
Plant Physiol., 95:450-455 (1991)
Characterization of adaptation in phototropism of Arabidopsis thaliana, Janoudi,A.-K., Poff,K.L. Plant Physiol.,
95: 517-521 (1991)

UV-B-inducible and temperature-sensitive photoreactivation of cyclobutane pyrimidine dimers in Arabidopsis


thaliana, Pang,Q., Hays,J.B. Plant Physiol., 95:536-543 (1991)

Substrate regulation of single potassium and chloride ion channels in Arabidopsis plasma membrane, Lew,R.R.
Plant Physiol., 95:642-647 (1991)

A Brassica S locus gene promoter directs sporophytic expression in the anther tapetum of transgenic Arabidopsis,
Toriyama, K., Thorsness,M.K., Nasrallah,J.B., Nasrallah,M.E. Dev. Biol., 143:427-431 (1991)

Arabidopsis Sensitivity of growth to high temperature, Binelli, G., Mascarenhas,J.P. Dev. Genet., 11:294-298
(1990)

The mutagenicity of azido derivatives of monosaccharides and alcohols and of N-(3-azido-2-hydroxypropyl)


derivatives of purines and pyrimidines in Arabidopsis thaliana and in Salmonella typhimurium, Gichner,T.,
Holy,A., Spassova,M., Veleminsky, J., Gruz,P. Mutagenesis, 6:55-58 (1991)

Status report of the International Programme on Chemical Safety's Collaborative Study on plant test systems,
Sandhu,S.S., De Serres,F.J., Gopalan,H.N.B., Grant,W.F., Veleminsky,J., Becking, G.C. Mut. Res. 257: 19-25
(1991)

Transgenic Arabidopsis thaliana plants obtained by particle bombardment mediated transformation, Seki,M.,
Shigemoto,N., Komeda,Y., Imamura,J., Morikawa,H. Appl. Microbiol. Biotechnol. 26:228-230 (1991)

Identification and map position of YAC clones comprising one third of the Arabidopsis genome, Hwant,I.,
Kohchi,T., Huauge,B., Goddman,H., Schmidt,R., Cnops,G., Dean,C., Gibson, S., Iba,K., Lemieux,B., Arondel,V.,
Danhoff,L., Somerville,C.R. Plant J. 1:367-374 (1991)

Plant lipids: mutants, metabnolism and membranes, Somerville,C.R., Browse,J. Science 252:80-87 (1991)

Chromosome walking in Arabidopsis using yast artificial chromosomes, Grill,E., Somrville,C.R. Molec. Gen.

37
Genet. 226:484-490 (1991)

38
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